<p>The distribution of plant species and functional traits in alpine and arctic environments are determined by abiotic conditions, but also by biotic interactions. In this thesis, I investigate interactions among plants and herbivory effects on plant community composition and plant functional traits in three different regions: Swedish Lapland, Beringia (USA/Russia) and Finnmark (Norway). Reindeer grazing was found to be extensive in southern Lapland and had limited effects on plant community composition and seedling germination. However, reindeer presence was found to influence plant functional traits, particularly in the subalpine birch forest. Tall herbs were lower and had lower SLA when reindeer were present, while small herbs showed an opposite pattern. The contrasting effects on the two herb groups are probably explained by a competitive release for small herbs when the tall herbs are suppressed by reindeer. Rodents had the largest relative impact on plant community composition in southern Lapland and this is consistent with the study from Finnmark, where rodents heavily affected dwarf shrubs on predator-free islands. With no predators present, vole densities increased profoundly and almost depleted some dwarf shrub species. These results support the idea that small mammals in arctic and alpine tundra are controlled by predators (i.e. top-down). However, a decrease in the nutritional quality in a sedge after defoliation gives support for the idea that small mammals are regulated by plant quality (i.e. bottom-up). In Beringia, small and large herbivores differed in the relation to plant community composition, since large herbivores were related to species richness and small herbivores were related to plant abundance. Plant functional traits were related only to large herbivores and standing crop of vascular plants.</p>

<p>Tolerance to herbivory is the ability of plants to maintain fitness in spite of damage. The goal of this thesis is to investigate the genetic variation and expression of tolerance within species, determine whether and in what conditions tolerance has negative side-effects, and how tolerance is affected by different ecological factors. Tolerance is investigated with special focus on the effects of different damage types, competitive regimes, history of herbivory, and polyploidization in plants. Studies are conducted as a literature review and three experiments on two cruciferous species Raphanus raphanistrum and Cardamine pratensis.</p><p>In the tolerance experiments, plants are subjected to artificial damage solely, or in a combination with natural damage. A literature review was conducted in order to investigate the effects of damage method. We found that traits related to tolerance, such as growth and fitness were not as sensitive in regard to damage method as measures of induced chemical traits, or measures of secondary herbivory.</p><p>Genetic variation of tolerance was demonstrated within populations of R. raphanistrum and between subspecies of C. pratensis. In R. raphanistrum, traits involved in floral display and male fitness were positively associated with plant tolerance to herbivore damage. A potential cost of tolerance was demonstrated as a negative correlation between levels of tolerance in high and low competitive regimes. I found no evidence of other proposed costs of tolerance in terms of highly tolerant plants suffering of reduced fitness in the absence of herbivores or trade-offs in terms of a negative association between tolerance to apical and leaf damage, or between tolerance and competitive ability. In C. pratensis, higher ploidy level in plants involved higher levels of tolerance measured as clonal reproduction. Furthermore, populations exposed to higher levels of herbivory had better tolerance than populations exposed to lower levels of herbivory. In this thesis, I demonstrate evidence of different components for the evolution of tolerance in plants: genotypic variation, selective factors in terms of costs and ploidization, and selective agents in terms of changing environment or herbivore pressure.</p>

<p>I have, in this thesis, studied the interactions between gray-sided voles (Clethrionomys rufocanus) and tundra vegetation, on islands in, and mainland sites close to the lake Iešjávri, in northern Norway. As isolated islands are virtually free of predation, I have been able to compare plant-herbivore interactions in the presence and absence of predators. I transplanted vegetation from an island with predators and voles, to predator-free islands with and with out voles. The results reveal the existence of a terrestrial trophic cascade as voles had a severe impact on the transplanted vegetation on the predator-free islands, but only minor effects on the mainland where predators are present. Moreover, this study shows that plant defence was only a successful strategy when predators were present. Voles reduced the abundance of all available plants during winter on the predator-free islands. The results imply that cascading effects of predation are most important for well-defended plants with grazing-sensitive morphology as these plants escape herbivore impacts in the presence of predators but are vulnerable in their absence</p><p>I studied the recovery of intensively grazed vegetation by building exclosures on islands that have been heavily grazed by voles for almost a decade.This study shows that the collective biomass of vascular plants recovered completely on three years, when voles were excluded. Although most species that are dominating the ungrazed vegetation recovered rapidly in the exclosures, the vegetation did not simply return to its ungrazed state. Herbaceous plants increased and there were pronounced differences in response among evergreen dwarf-shrub species. The semi-prostrate and tannin rich crowberry (Empetrum nigrum), showed the strongest recovery of all species, while the erect lingonberry (Vaccinium vitis-idaea) only showed weak signs of recovery. Thus, growth form determined the potential of plants to recover, whereas the trade-off between defensive investments and capacity to recover was weak or absent.</p><p>I studied the interaction between gray-sided voles and their main winter food plant, bilberry (Vaccinium myrtillus) on islands in and mainland sites close to the lake Iešjávri. I compared the abundance, population structure and palatability of bilberry ramets between vole-free islands, islands with voles but no predators and mainland sites with both voles and predators. Voles decreased the abundance of bilberry on the mainland, but the effect was much stronger on predator-free islands. Bilberry was fairly tolerant to grazing as it partially compensated for the lost tissue by producing more new ramets. Moreover, a cafeteria experiment showed that voles preferred the ramets from predator-free islands, which is inconsistent with conjectures emphasizing inducible plant defenses. The vole-bilberry interaction lacks features of delayed density dependence that could explain the vole cycles. I conducted a clipping and fertilization experiment to further investigate the effects of herbivory on palatability of bilberry shoots. Fertilization decreased the concentration of condensed tannins in shoots of bilberry and voles preferred fertilized and clipped shoots. I found no indication of induced defense that could reduce the palatability of bilberry twigs in response to herbivory.</p><p>The relationships between gray-sided vole densities, levels of invertebrate herbivory and chemical quality of leaves of Northern willow (Salix glauca) were studied on islands and mainland sites with contrasting vole densities. I found a positive correlation between level of invertebrate herbivory and vole density. The number of leaves per shoot, leaf size and leaf nitrogen content were also positively correlated with vole densities, while leaf C/N ratios were negatively correlated with vole densities. The positive correlation between vole densities and level of invertebrate herbivory is probably due to a facilitative effect of voles on invertebrate herbivores, mediated through changes in plant chemistry.</p>

I have, in this thesis, studied the interactions between gray-sided voles (Clethrionomys rufocanus) and tundra vegetation, on islands in, and mainland sites close to the lake Iešjávri, in northern Norway. As isolated islands are virtually free of predation, I have been able to compare plant-herbivore interactions in the presence and absence of predators. I transplanted vegetation from an island with predators and voles, to predator-free islands with and with out voles. The results reveal the existence of a terrestrial trophic cascade as voles had a severe impact on the transplanted vegetation on the predator-free islands, but only minor effects on the mainland where predators are present. Moreover, this study shows that plant defence was only a successful strategy when predators were present. Voles reduced the abundance of all available plants during winter on the predator-free islands. The results imply that cascading effects of predation are most important for well-defended plants with grazing-sensitive morphology as these plants escape herbivore impacts in the presence of predators but are vulnerable in their absence I studied the recovery of intensively grazed vegetation by building exclosures on islands that have been heavily grazed by voles for almost a decade.This study shows that the collective biomass of vascular plants recovered completely on three years, when voles were excluded. Although most species that are dominating the ungrazed vegetation recovered rapidly in the exclosures, the vegetation did not simply return to its ungrazed state. Herbaceous plants increased and there were pronounced differences in response among evergreen dwarf-shrub species. The semi-prostrate and tannin rich crowberry (Empetrum nigrum), showed the strongest recovery of all species, while the erect lingonberry (Vaccinium vitis-idaea) only showed weak signs of recovery. Thus, growth form determined the potential of plants to recover, whereas the trade-off between defensive investments and capacity to recover was weak or absent. I studied the interaction between gray-sided voles and their main winter food plant, bilberry (Vaccinium myrtillus) on islands in and mainland sites close to the lake Iešjávri. I compared the abundance, population structure and palatability of bilberry ramets between vole-free islands, islands with voles but no predators and mainland sites with both voles and predators. Voles decreased the abundance of bilberry on the mainland, but the effect was much stronger on predator-free islands. Bilberry was fairly tolerant to grazing as it partially compensated for the lost tissue by producing more new ramets. Moreover, a cafeteria experiment showed that voles preferred the ramets from predator-free islands, which is inconsistent with conjectures emphasizing inducible plant defenses. The vole-bilberry interaction lacks features of delayed density dependence that could explain the vole cycles. I conducted a clipping and fertilization experiment to further investigate the effects of herbivory on palatability of bilberry shoots. Fertilization decreased the concentration of condensed tannins in shoots of bilberry and voles preferred fertilized and clipped shoots. I found no indication of induced defense that could reduce the palatability of bilberry twigs in response to herbivory. The relationships between gray-sided vole densities, levels of invertebrate herbivory and chemical quality of leaves of Northern willow (Salix glauca) were studied on islands and mainland sites with contrasting vole densities. I found a positive correlation between level of invertebrate herbivory and vole density. The number of leaves per shoot, leaf size and leaf nitrogen content were also positively correlated with vole densities, while leaf C/N ratios were negatively correlated with vole densities. The positive correlation between vole densities and level of invertebrate herbivory is probably due to a facilitative effect of voles on invertebrate herbivores, mediated through changes in plant chemistry.

The distribution of plant species and functional traits in alpine and arctic environments are determined by abiotic conditions, but also by biotic interactions. In this thesis, I investigate interactions among plants and herbivory effects on plant community composition and plant functional traits in three different regions: Swedish Lapland, Beringia (USA/Russia) and Finnmark (Norway). Reindeer grazing was found to be extensive in southern Lapland and had limited effects on plant community composition and seedling germination. However, reindeer presence was found to influence plant functional traits, particularly in the subalpine birch forest. Tall herbs were lower and had lower SLA when reindeer were present, while small herbs showed an opposite pattern. The contrasting effects on the two herb groups are probably explained by a competitive release for small herbs when the tall herbs are suppressed by reindeer. Rodents had the largest relative impact on plant community composition in southern Lapland and this is consistent with the study from Finnmark, where rodents heavily affected dwarf shrubs on predator-free islands. With no predators present, vole densities increased profoundly and almost depleted some dwarf shrub species. These results support the idea that small mammals in arctic and alpine tundra are controlled by predators (i.e. top-down). However, a decrease in the nutritional quality in a sedge after defoliation gives support for the idea that small mammals are regulated by plant quality (i.e. bottom-up). In Beringia, small and large herbivores differed in the relation to plant community composition, since large herbivores were related to species richness and small herbivores were related to plant abundance. Plant functional traits were related only to large herbivores and standing crop of vascular plants.

Tolerance to herbivory is the ability of plants to maintain fitness in spite of damage. The goal of this thesis is to investigate the genetic variation and expression of tolerance within species, determine whether and in what conditions tolerance has negative side-effects, and how tolerance is affected by different ecological factors. Tolerance is investigated with special focus on the effects of different damage types, competitive regimes, history of herbivory, and polyploidization in plants. Studies are conducted as a literature review and three experiments on two cruciferous species Raphanus raphanistrum and Cardamine pratensis. In the tolerance experiments, plants are subjected to artificial damage solely, or in a combination with natural damage. A literature review was conducted in order to investigate the effects of damage method. We found that traits related to tolerance, such as growth and fitness were not as sensitive in regard to damage method as measures of induced chemical traits, or measures of secondary herbivory. Genetic variation of tolerance was demonstrated within populations of R. raphanistrum and between subspecies of C. pratensis. In R. raphanistrum, traits involved in floral display and male fitness were positively associated with plant tolerance to herbivore damage. A potential cost of tolerance was demonstrated as a negative correlation between levels of tolerance in high and low competitive regimes. I found no evidence of other proposed costs of tolerance in terms of highly tolerant plants suffering of reduced fitness in the absence of herbivores or trade-offs in terms of a negative association between tolerance to apical and leaf damage, or between tolerance and competitive ability. In C. pratensis, higher ploidy level in plants involved higher levels of tolerance measured as clonal reproduction. Furthermore, populations exposed to higher levels of herbivory had better tolerance than populations exposed to lower levels of herbivory. In this thesis, I demonstrate evidence of different components for the evolution of tolerance in plants: genotypic variation, selective factors in terms of costs and ploidization, and selective agents in terms of changing environment or herbivore pressure.

The aim of this thesis project was to elucidate patterns and processes associated with the biotic interactions in a natural plant–herbivore–parasitoid food web characterized by spatial and temporal heterogeneity with regard to species composition. The system examined is based on island populations of the perennial herb Meadowsweet (Filipendula ulmaria, Rosaceae), located in the Skeppsvik Archipelago. The area is subject to isostatic rebound, amounting to 0.85 cm per year; this makes it possible to calculate the age of the rising islands. Meadowsweet colonizes new islands when they are about 100 years old. Meadowsweet is consumed by two major herbivores in the study area: Galerucella tenella and Altica engstroemi (Coleoptera: Chrysomelidae). Both herbivores overwinter in the topsoil and successful colonization occurs when the islands reach a height that prevents the beetles from being removed or killed as a result of wave wash during the winter. I found that both herbivores significantly reduced individual plant fitness and population growth rate. A “cafeteria experiment” with Galerucella showed that this beetle discriminated between plants from different islands, avoiding plants from old islands which contained high concentrations of putative defence compounds, while readily accepting plants from younger islands which contained lower concentrations of these chemicals. Further, the plant species exhibited a trade-off between growth and production of the putative defence compounds. Taken together, these results were interpreted as providing evidence of herbivore-driven evolution of resistance in Meadowsweet. Further, laboratory studies suggested that Galerucella gradually includes a less preferred host plant (Rubus arcticus, Rosaceae) in its diet as Meadowsweet resistance increases. This implies that Galerucella drives its own host-breadth enlargement by selectively inducing a ‘rent rise’ in the original host, Meadowsweet. In a number of field studies I showed that the oligophagous parasitoid Asecodes mento (Hymenoptera: Eulophidae) has a strong positive effect on Meadowsweet seed set by removing large numbers of G. tenella larvae. This top-down effect is, however, altered by the presence of a close relative of G. tenella, namely G. calmariensis, which is monophagous on Purple loosestrife (Lythrum salicaria, Lythraceae). G. tenella experiences associational susceptibility when coexisting with G. calmariensis since the latter supports a higher and more fit pool of shared parasitoids and because Meadowsweet attracts a higher proportion of the shared parasitoid females than Purple loosestrife. This implies that G. tenella densities are very low in coexisting populations and that Meadowsweet experiences associational resistance and produces more seeds when co-occurring with Purple loosestrife. Thus, selection for increased resistance in Meadowsweet is likely to be relaxed in populations mixed with Purple loosestrife. I conclude that the evolution of plant resistance is likely to depend on the length of time and intensity of selection. When Meadowsweet colonizes new islands it experiences a period of enemy-free space; followed by a midlife and ageing with selection by herbivores. The intensity of this selection does, however, depend on the presence of additional plant and herbivore species.

Chemically mediated interactions have important ecological and evolutionary effects on populations and communities. Despite recognition that herbivory can significantly affect the biomass and composition of freshwater macrophyte communities, there are few investigations of chemical defenses among freshwater vascular plants and mosses and none of freshwater red algae. This study compares the palatability of five species of freshwater red algae (Batrachospermum helminthosum, Boldia erythrosiphon, Kumanoa sp., Paralemanea annulata, and Tuomeya americana) that occur in the southeastern United States relative to two co-occurring macrophytes (the chemically defended aquatic moss Fontinalis novae-angliae and the broadly palatable green alga Cladophora glomerata). We assessed the potential role of structural, nutritional, and chemical traits in reducing macrophyte susceptibility to generalist crayfish grazers. Both native and non-native crayfish significantly preferred the green alga C. glomerata over four of the five species of red algae. B. erythrosiphon was palatable, while the cartilaginous structure of P. annulata reduced its susceptibility to grazing, and chemical defenses of B. helminthosum, Kumanoa sp., and T. americana rendered these species as unpalatable as the moss F. novae-angliae. Extracts from these latter species reduced feeding by ~30-60% relative to solvent controls if tissues were crushed (simulating herbivore damage) prior to extraction in organic solvents. However, if algae were first soaked in organic solvents that inhibit enzymatic activity and then crushed, crude extracts stimulated or had no effect on herbivory. B. helminthosum, Kumanoa sp., and T. americana all exhibited "activated" chemical defenses in which anti-herbivore compounds are produced rapidly upon herbivore attack via enzymatic processes. In an additional accept/reject behavioral assay, B. helminthosum extracts reduced the number of crayfish willing to feed by >90%. Given that three of the five red algal taxa examined in this study yielded deterrent crude extracts, selection for defensive chemistry in freshwater rhodophytes appears to be substantial. Activated chemical defenses are thought to be an adaptation to reduce the resource allocation and ecological costs of defense. As such, activated chemical defenses may be favored in freshwater red algae, whose short-lived gametophytes must grow and reproduce rapidly. Roughly 20% of the known chemical defenses produced by marine algae are activated; further examination is needed to determine whether the frequency of activated chemistry is higher in freshwater red algae compared to their marine counterparts. Continued investigation of chemical defenses in freshwater red algae will contribute to among-system comparisons, providing new insights in the generality of plant-herbivore interactions and their evolution.

The sacoglossan sea slug Elysia clarki is able to photosynthesize for three to four months using chloroplasts sequestered from its algal food sources. Furthermore, the slug is able to store multiple chloroplasts from different algal species within the same cell. This research, consisting of several related studies, explores the role that provision of organic nutrients via photosynthesis plays in the biology of the slug. The first chapter demonstrates that, under conditions of starvation, photosynthetic activity in E. clarki remains fully functional for one month after which it then declines. During the first month of starvation the slug exhibits similar feeding behavior as slugs provided a continuous supply of food, suggesting that photosynthesis delays the onset of starvation-induced behavioral changes. The second chapter explores E. clarki's spatial relationships with algae known to be food sources in the field. In areas with high slug density, edible algal populations were very low. DNA barcoding was employed to demonstrate that the algae found near slugs were poor predictors of which foods were actually consumed by slugs. Generally, there was a mismatch between algae available in the field and slug diets. The third chapter explores how E. clarki is able to maintain photosynthesis. After labeling with a C14 ALA incubation process, then chlorophyll was extracted from slugs and purified using HPLC. Results indicate that recently collected E. clarki are able to synthesize chlorophyll, whereas slugs starved for 3 months were not. Photosynthesis plays a very important role for E. clarki and its relationships with food algae.

Using a comparative approach, the presented study explores the ecology of ten species of native New Zealand broom, Carmichaelia, and their vulnerability to competition and herbivory, with the aim of gaining a better understanding of the significance of introduced species as a threat to rare indigenous plants in New Zealand. In particular, the study focuses on the relationship between characteristics of the Carmichaelia species and their vulnerability, as well as on other factors influencing the significance of introduced species as a threat. To gain a better understanding of the ecology of the Carmichaelia species, their current habitats and associated plant communities were investigated using quantitative-descriptive methods in the field. The effect of competition with introduced plants was studied in two glasshouse experiments, differentiating above ground competition for light from below ground competition for nutrients and water. The experiments focussed on the early life-stage of seedling establishment of the Carmichaelia species. The impact of herbivory by introduced mammals was studied in four field-based exclosure trials, focussing on the effects on survival and reproductive activity of adult Carmichaelia plants. The results showed that the effects of competition and herbivory vary between the different species. Furthermore, they provided a set of species characteristics that can be used as indicators to predict the vulnerability of Carmichaelia to the impact of introduced species. These indicators provide a useful tool for threatened species management, as they allow the identification of the most vulnerable species as well as the most significant threat to each species. Furthermore, the indicators can be used to group species, combining those with similar vulnerability profiles, and therefore, likely similar management needs. However, the example of the Carmichaelia species also illustrated that the use of indicators for the vulnerability of threatened species is limited and needs to be combined with case-by-case studies to verify the actual significance of threats for each population of concern. The vulnerability profiles derived from species’ characteristics can be used to guide such site specific studies, ensuring they focus on the most relevant threat factors. This combination of the understanding of general patterns in the vulnerability of species with targeted species and site-specific studies will lead to increased efficiency in the conservation management of threatened plant species.